The Ethiopian wolf has a very restricted range. It is found only in six or seven mountain ranges of Ethiopia. This includes the Arssi and Bale mountains of southeast Ethiopia, the Simien mountains, northeast Shoa, Gojjam, and Mt. Guna (Ginsberg and Macdonald 1990). The largest population exists in the Bale Mountains National Park with 120-160 individuals (Sillero-Zubiri and Gottelli 1995).
Biogeographic Regions: ethiopian (Native )
Endemic to the Ethiopian highlands, above the tree line at about 3,200 m. There are no recent records of the species at altitudes below 3,000 m, although specimens were collected at 2,500 m from Gojjam and north-western Shoa at the beginning of the century (references in Yalden et al. 1980). Reported in the Simien Mountains since the species was first described in 1835, but scattered and irregular sightings suggest numbers have been declining. Reported on the Gojjam plateau until the early 1900s (Powell-Cotton 1902, Maydon 1932). South of the Rift Valley, wolves have been reported in the Arsi Mountains since the 1920s, and, more recently (1959), in the Bale Mountains. Reports of small populations in North Sidamo (Haltenorth and Diller 1980) may be in error. There is no evidence that the Ethiopian Wolf ever occurred in Eritrea (Coetzee 1977).
The species currently is confined to seven isolated mountain ranges of the Ethiopian highlands, at altitudes of 3,000–4,500 m (Gottelli and Sillero-Zubiri 1992, Marino 2003a). In the northern highlands wolves are restricted to land above 3,500–3,800 m by increasing agricultural pressure (Yalden et al. 1980, Marino 2003a). Wolf populations occur north of the Rift Valley in the Simien Mountains, Mount Guna, North Wollo and South Wollo highlands, and Menz. Recently extinct in Gosh Meda (North Shoa), and absent from Mt Choke, Gojjam, for a few decades. South-east of the Rift Valley there are populations in the Arsi Mountains (Mt Kaka, Mt Chilalo and Galama range) and in the Bale Mountains, including the Somkaru-Korduro range (Marino 2003a).
Ethiopian wolves are long-limbed, slender looking canids. They have a reddish coat with white marking on the legs, underbelly, tail, face, and chin. The boundary between the red and white fur is quite distinct. White markings on the face include a characteristic white crescent below the eyes and a white spot on the cheeks. The chin and throat are also white. The tail is marked with an indistinct black stripe down its length and a brush of black hairs at the tip. The ears are wide and pointed and the nose, gums, and palate are black. Females are generally paler in color than males and are smaller overall. There are five toes on the front feet and four on the rear feet. Males measure from 928 to 1012 mm (average 963 mm) and females from 841 to 960 mm (average 919 mm). Males weigh from 14.2 to 19.3 kg (average 16.2) and females from 11.2 to 14.2 kg (average 12.8). The tail is from 270 to 396 mm in length. The dental formula is 3/3:1/1:4/4:2/3, with the lower third molar being absent occasionally. (Sillero-Zubiri and Marino, 1995)
Range mass: 11.2 to 19.3 kg.
Range length: 841 to 1012 mm.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: male larger
Canis simensis is found in afro-alpine grasslands and heathlands where vegetation is less than 0.25 m high. It lives at altitudes of 3000-4400 m (Sillero-Zubiri and Gottelli 1994).
Range elevation: 3000 to 4400 m.
Habitat Regions: temperate ; terrestrial
Terrestrial Biomes: savanna or grassland ; mountains
Habitat and Ecology
A very localized endemic species, confined to isolated pockets of Afroalpine grasslands and heathlands where they prey on Afroalpine rodents. Suitable habitats extend from above treeline at about 3,200 m up to 4,500 m, with some wolves present in montane grasslands at 3,000 m. However, subsistence agriculture extends up to 3,500–3,800 m in many areas, restricting wolves to higher ranges (Marino 2003a). Rainfall at high altitude varies between 1,000 and 2,000 mm/year, with one pronounced dry period from December to February/March.
Wolves utilize all Afroalpine habitats, but prefer open areas with short herbaceous and grassland communities where rodents are most abundant, along flat or gently sloping areas with deep soils and poor drainage in parts. Prime habitats in the Bale Mountains are characterized by short herbs (Alchemilla spp.) and grasses and low vegetation cover, a community maintained in continuous succession as a result of molerat (Tachyoryctes macrocephalus) burrowing activity. Other good habitats include tussock grasslands (Festuca spp., Agrostis spp.), high-altitude scrubs dominated by Helichrysum spp. and short grasslands in shallow soils. In northern parts of the range, plant communities characterized by a matrix of 'guassa' tussock grasses (Festuca spp.), 'cherenfi' bushes (Euryops pinifolius) and giant lobelias (Lobelia rhynchopetalum) sustain high rodent abundance and are preferred by wolves. Ericaceous moorlands (Erica and Phillipia spp.) at 3,200–3,600 m are of marginal value, with open moorlands having patches of herbs and grasses which are relatively good habitat.
Canis simensis is a carnivore, generally preying on rodents ranging in size from the giant mole-rat Tachyoryctes macrocephalus (900 g) to that of the common grass rats (Arvicanthis blicki, Lophuromys melanonyx; 90-120 g) (Ginsberg and Macdonald 1990). In 689 feces, murid rodents accounted for 95.8% of all prey items, and 86.6% belonged to the three species listed above (Sillero-Zubiri and Gottelli 1994). When present in the hunting range, giant mole-rats are the primary component of the diet. In its absence, the common mole-rat Tachyoryctes splendens is most commonly eaten (Malcom 1997). Canis simensis also eats goslings, eggs, and young ungulates (reedbuck and mountain nyla) and occasionally scavenges carcasses. The Ethiopian wolf often caches its prey in shallow holes (Ginsberg and Macdonald 1990).
Prey is usually captured by digging it out of burrows. Areas of high prey density are patrolled by wolves walking slowly. Once prey is located, the wolf moves stealthily towards it and grabs it with its mouth after a short dash. Occasionally, the Ethiopian wolf hunts cooperatively to bring down young antelopes, lambs, and hares (Sillero-Zubiri and Gottelli 1994).
Animal Foods: birds; mammals; eggs; carrion
Foraging Behavior: stores or caches food
Primary Diet: carnivore (Eats terrestrial vertebrates)
Ethiopian wolves are top predators in the ecosystems in which they live.
Life History and Behavior
Perception Channels: tactile ; chemical
Ethiopian wolves may live 8 to 10 years in the wild, although one wild individual was recorded living to 12 years. (Sillero-Zubiri and Marino, 1995)
Status: wild: 12 (high) years.
Status: wild: 10 (high) years.
For Ethiopian wolves, dispersal from their native packs is limited due to habitat saturation. Males generally remain in their natal pack, and a small number of females disperse in their second or third year. To combat this high potential for inbreeding inside the closely related pack, matings outside the pack occur frequently. Copulation outside the pack occurs with males of all rank, but those within the pack occur only between the dominant male and female. While copulation between males and subordinate females does occur, pups that may arise from this union rarely survive (Sillero-Zubiri et al. 1996).
Prior to copulation, the dominant female increases her rate of scent marking, play soliciting, food begging towards the dominant male, and aggressive behavior towards subordinate females. Ethiopian wolves mate over a period of 3-5 days, involving a copulation tie that lasts up to 15 minutes.
It is not uncommon for a subordinate female to assist in suckling the young of the dominant female. In these cases, the subordinate lactating female is likely pregnant and either loses or deserts her own young for those of the dominant female.
Mating System: monogamous ; cooperative breeder
Once a year between October and January, the dominant female in each pack gives birth to a litter of 2-6 pups. Gestation lasts approximately 60-62 days. The female gives birth to her litter in a den she digs in open ground under a boulder or in a rocky crevice. The pups are born with their eyes closed and no teeth. They are charcoal gray with a buff patch on their chest and under areas. At about 3 weeks, the coat begins to be replaced by the normal adult coloring and the young first emerge from the den. After this time, den sites are regularly shifted, sometimes up to 1300m.
Development of the young occurs in three stages (Sillero-Zubiri and Gottelli 1994). The first covers weeks 1-4 when the pups are completely dependent on their mother for milk. The second occurs from week 5-10 from when the pups' milk diet is supplemented by solid food regurgitated from all pack members. It ends when the pups are completely weaned. Finally, from week 10 until about 6 months, the young survive almost solely on solid food provided from adult members of the pack. Adults have been seen providing food for young up to 1 year old. The Ethiopian wolf attains full adult appearance at 2 years of age, and both sexes are sexually mature during their second year (Sillero-Zubiri and Gottelli 1994). Data on life expectancy is inadequate, but C. simensis is likely to live 8-9 years in the wild (Macdonald 1984).
Range number of offspring: 2 to 6.
Range gestation period: 60 to 62 days.
Average weaning age: 70 days.
Key Reproductive Features: gonochoric/gonochoristic/dioecious (sexes separate); sexual
Parental Investment: altricial ; post-independence association with parents; extended period of juvenile learning
Ethiopian wolves are considered endangered by both the IUCN and U.S. Endangered Species Act. They are protected from hunting under Ethiopian law. Effort to curb the transmission of diseases, especially rabies, to Ethiopian wolves from domestic dogs and to prevent hybridization with domestic dogs have been undertaken. In addition, monitoring of Ethiopian wolf populations continues. (Sillero-Zubiri and Marino, 1995)
US Federal List: endangered
CITES: no special status
IUCN Red List of Threatened Species: endangered
IUCN Red List Assessment
Red List Category
Red List Criteria
Bale minimum estimate of N=210 adults was derived from intensive observations of wolf packs across the Bale Mountains. In April 2009 the population was estimated at 291 adult (>1 year old) wolves (EWCP annual report 2008–2009) and in April 2010 at 290–320 (EWCP annual report 2009-2010). Since then 23 carcasses have been retrieved, and 56 individuals have gone missing, probably as a result of a canine distemper epizootic that affected several subpopulations in Bale.
Ethiopian Wolves are restricted to seven isolated mountain enclaves in the highlands of Ethiopia. The Afroalpine range available to the species is shrinking and increasingly fragmented. The species’ extent of occurrence is estimated at 4,200 km² with an area of occupancy of 2,779 km². Population estimates across the species’ range suggest that between 360 and 440 adult (>1 year old) wolves remain, of which <250 are mature individuals. One relict population in northern Ethiopia (Mt Guna) is now functionally extinct.
The largest population occurs in the Bale Mountains in southern Ethiopia, where intensive monitoring of wolf packs indicate that population size varies widely due to the recurrence of infectious disease epizootics affecting local subpopulations. Since 2008 to date this population has declined by ~30%, from approximately 300 to an estimated 210 adults, due to consecutive epizootics of rabies and canine distemper. This equates to a 20% global decline in the population in the last three years; although narrowly below the threshold for uplisting to Critically Endangered under C1, this continued decline merits careful ongoing monitoring of population numbers (and a reassessment if necessary) and sustained conservation interventions.
Number of "Mature Individuals"
The number of mature individuals in the population (population size) was determined using the IUCN's new definition (in: Guidelines for Using the IUCN Red List Categories and Criteria) and the following formula:
Mature individuals = 0.31N1+ 0.4N1+0.57N2 +0.16 N2 then
Mature individuals = 0.71N1+ 0.73 N2 where:
N1= adult male population
N2= adult female population
Calculations of mature individuals capable of reproduction were derived from empirical data on wolf populations in the Bale Mountains, between 1988–1992 and 1997–2000. Of all adult males in a population, 31% are alpha males and thus reproductive individuals (= 0.31N1). Of all subordinate adult males (= 0.69N1), some may access reproduction either by replacing the alpha male or through extra-pack copulations. This proportion was subjectively estimated at 60% and thus the contribution of subordinate adult males potentially capable of reproduction is 0.6 x 0.69N1 = 0.41N1.
Of all adult females in a population, 57% are in alpha position and thus reproduce (= 0.57N2). Of the suppressed adult females (= 0.33N2) we estimated a 50% likelihood to access reproduction in their lifetime (0.5 x 0.33N2=0.16N2). This was derived from the rate of breeding opportunities, calculated as 0.12 for female per year from empirical data, over four years (0.12 x 4 years= 0.5).
Population estimates were derived from current availability of habitat types and extrapolations of the density of adult and subadult wolves to the area of each habitat type. Population estimates (N) are numbers of adult and subadult wolves. Given a population sex ratio of 2:1 males to females then the male population is 2/3N = 0.67N and the female population is 1/3N= 0.34N. Eighty percent of males and 70% of females are adults, therefore:
N1= adult male population= 0.8 x 0.67N = 0.53N
N2= adult female population= 0.7 x 0.33N = 0.23N
and, Mature individuals = 0.71N1 + 0.73N2, so:
Mature individuals = 0.37N + 0.17N= 0.54N
Mature individual estimates for 2010 based on IUCN/SSC Canid Specialist Group (2011):
Bale Mountains: N = 210; mature individuals = 113
Arsi Mountains: N = 54; mature individuals = 29
Simien Mountains: N = 52; mature individuals = 28
North Wollo: N = 19; mature individuals = 10
South Wollo: N = 16; mature individuals = 9
Menz : N = 15; mature individuals = 8
Mt Guna: N = 0; mature individuals = 0
Total: N = 366; mature individuals = 197
Guassa-Menz estimate of 15 adult wolves was derived from intensive observation of wolf packs in recent years which yielded an estimate of 25 adult wolves, and was corroborated by DNA fingerprinting in 2008 (which identified 23 individuals). During 2010 40% of the known wolves have died or disappeared due to canine distemper, resulting in the revised estimate.
Arsi and Simien estimates (EWCP unpublished) were derived from intensive observations of wolf packs across both ranges. Recent improvements in monitoring resulted in a larger estimate for Simien, and a smaller one for Arsi. These revised figures reflect improved field techniques and effort rather than actual population change.
For North and South Wollo, population estimates are conservative and were based on habitat availability. Using field maps, the extent of suitable habitat were estimated and populations extrapolated from known densities in Bale for habitats with similar characteristics (Marino 2003a). Here the lower value of densities observed were used (i.e., good habitat: 0.2 wolves/km²; marginal habitat 0.1 wolf/km²).
Other parameters used:
Generation time: mean 4.5 yr, range 3–8; extent of occurrence: 4,200 km² [defined as area above 3,000 m asl]; area of occupancy: 2,800 km²; number of populations: 7; number of sub-populations: 10.
Given the information above, Canis simensis meets the criteria for Endangered under criterion B based on <5,000 km² extent of occurrence (B1), as the population is severely fragmented (a), and there is a continuing decline (b) in area, extent and or quality of habitat (iii) and number of mature individuals (v). It also meets the criteria under C based on <2,500 mature individuals in the population (EN C and D), exceeding a 20% decline in the population in the past eight years (two generations), and a continuing decline in population size (EN C2) and <250 mature individuals in each subpopulation (EN C2a(i)). Finally, it also meets the threshold of <250 mature individuals for listing under D.
- 1996Critically Endangered
- 1994Endangered(Groombridge 1994)
- 1990Endangered(IUCN 1990)
- 1988Endangered(IUCN Conservation Monitoring Centre 1988)
- 1986Endangered(IUCN Conservation Monitoring Centre 1986)
Date Listed: 07/27/1979
Lead Region: Foreign (Region 10)
Where Listed: Ethiopia
Population location: Ethiopia
Listing status: E
For most current information and documents related to the conservation status and management of Canis simensis , see its USFWS Species Profile
More than half of the species' population lives in the Bale Mountains, where wolf density is high for a social carnivore of its size, and positively correlated with density of rodent prey and negatively with vegetation height (Sillero-Zubiri and Gottelli 1995). Highest wolf densities are found in short Afroalpine herbaceous communities (1.0–1.2 adults/km²); lower densities are found in Helichrysum dwarf-scrub (0.2/km²), and in ericaceous heathlands and barren peaks (0.1/km²). Wolves are also present at low density (0.1–0.2/km²) in montane grasslands at lower altitudes.
Elsewhere, overall wolf density is relatively lower. In Menz, wolf density was estimated at 0.2 animals/km² using transect data (Ashenafi 2001). Comparison of census transect data from recent comprehensive surveys (Marino 2003a) indicates higher abundance in North Wollo (0.20 ± 0.20 sightings per km), intermediate in Arsi and Guna (0.10 ± 0.11 and 0.10 ± 0.14, respectively), and lower in South Wollo and Simien (0.08 ± 0.13 and 0.06 ± 0.11, respectively). These results were supported by counts of wolf signs (i.e., diggings and droppings) and interview results.
A summary of population abundance and status in each region can be found in Sillero-Zubiri et al. (2004), with the most recent information on population status found in IUCN/SSC Canid Specialist Group (2011), and summarized above. Time series from long-term monitoring of wolves in the Bale Mountains of southern Ethiopia, spanning over 20 years, indicated marked variation in wolf abundance in association to rabies epizootics affecting high-density populations in 1991, 2003 and 2008 (Marino 2003b, Marino et al. 2006, IUCN/SSC Canid Specialist Group 2011). Population numbers returned to previous levels after the first two disruptions, evidencing resilience to catastrophes, but at very low density population growth was unexpectedly low, due to delays in the formation of new breeding packs (Marino et al. in review). Canine distemper (CDV) appears to be more important than previously thought, with CDV outbreaks affecting wolves across subpopulations in the Bale Mountains in 2005, 2006 (Malcolm 2006), and 2010 (Gordon et al. 2010).
Continuous loss of habitat due to high-altitude subsistence agriculture represents the major threat. Sixty percent of all land above 3,200 m has been converted into farmland, and all populations below 3,700 m are particularly vulnerable to further habitat loss, especially if the areas are small and of relatively flat relief (Marino 2003a). Habitat loss is exacerbated by overgrazing of highland pastures by domestic livestock, and in some areas habitat is threatened by proposed development of commercial sheep farms and roads. Human persecution triggered by political instability in the past is currently less severe and associated with conflicts over livestock losses (Marino 2003a). Recent population decline in Bale has been mostly due to disease epizootics, with road kills and shooting as secondary threats. Rabies is a potential threat to all populations, with canine distemper also a concern in Bale. Most of these threats are exacerbated by the wolves' specialization to life in the Afroalpine ecosystem.
In Bale, the Ethiopian wolf hybridizes with domestic dogs (Canis familiaris). Gottelli et al. (1994) used mitochondrial DNA restriction fragments and microsatellite alleles to conclude that hybridization was relatively common in western Bale as a result of crosses between female wolves and male domestic dogs. Hybrids have shorter muzzles, heavier-built bodies and different coat patterns. Although hybrids are confined to the Web Valley in western Bale they may threaten the genetic integrity of the wolf population. Following hybridization, a population may be affected by outbreeding depression or reduction in fitness, although to date this does not seem to have taken place in Bale. Currently there is no indication of hybridization taking place outside western Bale.
There is no exploitation of the species for furs, body parts or other purposes.
The species is not included on the CITES Appendices. It has full official protection under Ethiopia's Wildlife Conservation Regulations of 1974, Schedule VI. Killing a wolf carries a sentence of up to two years.
The Ethiopian Wolf occurs in several protected areas: Bale Mountains National Park; Simien Mountains National Park; Borena Saiynt Regional Park (South Wollo); Guassa Community Conservation Area (North Shoa); Arsi Mountains Regional Park. As a result of boundary extensions (Simien) and new parks created (Arsi) the area of suitable wolf habitat within protected areas increased to 87%.
A number of important steps have been taken in the interests of conserving this endemic species, including: 1) a dog vaccination campaign in Bale, and vaccination campaigns in response to rabies outbreaks in dogs close to wolves in Menz and Simien; 2) sterilization programme for hybrids in Bale; 3) vaccination of wolves in response to rabies epizootics in parts of Bale; 4) community and school education programme in Bale and Wollo; 5) strengthening the capacity of the Bale Mountains National Park – funding patrolling, maintenance of infrastructure, etc.; 6) surveys to determine the persistence and status of all populations of wolves; 7) monitoring of all wolf populations; 8) A Strategic Planning workshop for Ethiopian wolf conservation held in Lalibela, February 2011, resulting in a National Action Plan for the next 10 years (IUCN/SSC Canid specialist Group 2011).
In 1983, the Wildlife Conservation Society established the Bale Mountains Research Project, which publicized the wolf's plight and started a regular monitoring programme for the species. A detailed four-year field study followed (Sillero-Zubiri 1994). Based on its findings, the IUCN/SSC Canid Specialist Group produced an action plan for the Ethiopian Wolf (Sillero-Zubiri and Macdonald 1997), which provided a detailed strategy for the conservation and management of remaining wolf populations. This plan advocated immediate action on three fronts – education, wolf population monitoring, and rabies control in domestic dogs – to conserve the Afroalpine ecosystem and its top predator. As a result, the Ethiopian Wolf Conservation Programme (EWCP) was established in 1995 by Oxford University with financial support from the Born Free Foundation, Frankfurt Zoological Society (FZS), the Wildlife Conservation Network and other donors. The Programme’s overall aim is to protect the Afroalpine ecosystem and many of its rare highland endemic plants and animals through better management in Bale and the establishment of other conservation areas in Menz and Wollo. The EWCP currently monitors the demography of the Bale and selected populations in South and North Wollo, supports park patrols within the wolf range, undertakes domestic dog control and the removal of dog-wolf hybrids. Additionally, the EWCP carries out a community conservation education campaign that targets people living inside the wolf's range and is aimed at improving dog husbandry and combating disease in the park and surroundings. A large-scale dog vaccination programme (targeting up to 5,000 dogs a year) seeks to reduce the occurrence of rabies and distemper within the Ethiopian wolf range and is backed up by further epidemiological and demographic studies. The EWCP is also active elsewhere in Ethiopia, with representatives surveying and monitoring all wolf ranges and implementing education campaigns about the plight of the species. Zelealem Tefera Ashenafi set up the Guassa Biodiversity Project in 1996, looking at the relationships between pastoralists and wildlife in the highlands of Menz. FZS and partners are working in North Ethiopia to protect the Afrolapine ecosystem, supporting the Simien Mountains National Park and creation of a new community conservation area in North Wollo.There are no animals maintained in captivity.
Although the behavioural ecology of the species is well known, research has focused largely in the optimal habitats in the Bale Mountains. Additional information on dispersal distance and survival would be useful. Investigation into the role of the species in the epidemiology of canid-related diseases is necessary. Studies on wolf-prey relationships and prey availability in the high risk populations of northern Ethiopia are also needed.
Relevance to Humans and Ecosystems
The Ethiopian wolf occasionally preys on lambs (Sillero-Zubiri 1995).
Canis simensis helps control populations of rodents in its habitat.
The Ethiopian wolf (Canis simensis) is a canid native to the Ethiopian Highlands. It is similar to the coyote in size and build, and is distinguished by its long and narrow skull, and its red and white fur. Unlike most large canids, which are widespread, generalist feeders, the Ethiopian wolf is a highly specialised feeder of Afroalpine rodents with very specific habitat requirements. It is one of the world's rarest canids, and Africa's most endangered carnivore.
The species' current range is limited to seven isolated mountain ranges at altitudes of 3,000–4,500m, with the overall adult population estimated at 360-440 individuals in 2011, more than half of them in the Bale Mountains.
The Ethiopian wolf is listed as Endangered by the IUCN, on account of its small numbers and fragmented range. Threats include increasing pressure from expanding human populations, resulting in habitat degradation through overgrazing and disease transference from free ranging dogs. Its conservation is headed by Oxford University's Ethiopian Wolf Conservation Programme (EWCP), which seeks to protect wolves through vaccination and community outreach programs.
- 1 Naming
- 2 Historical account
- 3 Taxonomy and evolution
- 4 Physical description
- 5 Behaviour
- 6 Ecology
- 7 Range and populations
- 8 Threats
- 9 Conservation
- 10 Notes
- 11 References
- 12 Bibliography
- 13 Further reading
- 14 External links
|Linguistic group or area||Indigenous name||Literal translation|
|Amharic||ቀይ ቀበሮ (Ky kebero)|
|Lupos Ethiopia mittit, cervice iubatos et tanto varios ut nullum eis colorem dicunt abesse. Ethiopicis lupis proprium est, quod in saliendo ita nisus habent alitis, ut non magis proficient cursu quam meatu. Homines tamen numquam impetunt. Bruma comati sunt, aestate nudi. Ethiopes eos vocant theas.||Ethiopia produces wolves with manes, so diversely coloured, men say, that no hue is lacking. A characteristic of Ethiopian wolves is that they leap so high that they seem to have wings, going further than they would by running. They never attack men, however. In winter, they grow long hair; in summer, they are hairless. The Ethiopians call them theas.|
The species was first scientifically described in 1835 by Eduard Rüppell, who provided a skull for the British Museum. European writers traveling in Ethiopia during the mid-19th century (then called Abyssinia) wrote that the animal's skin was never worn by natives, as it was popularly believed that the wearer would die should any wolf hairs enter an open wound, while Charles Darwin hypothesised that the species gave rise to greyhounds.[c] Since then, it was scarcely heard of in Europe up until the early 20th century, when several skins were shipped to England by Major Percy Horace Gordon Powell-Cotton during his travels in Abyssinia.
The Ethiopian wolf was recognised as requiring protection in 1938, and received it in 1974. The first in-depth studies on the species occurred in the 1980s, with the onset of the American sponsored Bale Mountains Research Project. Ethiopian wolf populations in the Bale Mountains National Park were negatively affected by the political unrest of the Ethiopian Civil War, though the critical state of the species was revealed during the early 1990s after a combination of shooting and a severe rabies epidemic decimated most packs studied in the Web Valley and Sanetti Plateau. In response, the IUCN reclassified the species from Endangered to Critically Endangered in 1994. The IUCN/SSC Canid Specialist Group advocated a three front strategy of education, wolf population monitoring, and rabies control in domestic dogs. The establishment of the Ethiopian Wolf Conservation Programme in Bale soon followed in 1995 by Oxford University, in conjunction with the Ethiopian Wildlife Conservation Authority (EWCA).
Soon after, a further wolf population was discovered in the Central Highlands. Elsewhere, information on Ethiopian wolves remained scarce; although first described in 1835 as living in the Simien Mountains, the paucity of information stemming from that area indicated that the species was likely declining there, while reports from the Gojjam plateau were a century out of date. Wolves were recorded in the Arsi Mountains since the early 20th century, and in the Bale Mountains in the late 1950s. The status of the Ethiopian wolf was re-assessed in the late 1990s, following improvements in travel conditions into northern Ethiopia. The surveys taken revealed local extinctions in Mount Choqa, Gojjam, and in every northern Afroalpine region where agriculture is well developed and human pressure acute. This revelation stressed the importance of the Bale Mountains wolf populations for the species' long-term survival, as well as the need to protect other surviving populations. A decade after the rabies outbreak, the Bale populations had fully recovered to pre-epizootic levels, prompting the species' downlisting to Endangered in 2004, though it still remains the world's rarest canid, and Africa's most endangered carnivore.
Taxonomy and evolution
The earliest fossil carnivores that can be linked with some certainty to canids are the Eocene Miacids, which lived some 38 to 56 million years ago. The Miacids later diverged into caniforms and feliforms, with the former line leading to such genera as the coyote-sized Mesocyon of the Oligocene (38 to 24 million years ago), the fox-like Leptocyon and the wolf-like Tomarctus which inhabited North America some 10 million years ago. Although there are fossil records of wolf-like canids from Late Pleistocene Eurasia, there are no fossil records for the Ethiopian wolf itself. The species may have evolved from a wolf-like ancestor crossing into North Africa from Eurasia as early as 100,000 years ago. Due to the high density of rodents in their new Afroalpine habitat, the ancestors of the Ethiopian wolf gradually developed into specialised rodent hunters. This specialisation is reflected in the animal's skull morphology, with its very elongated head, long jaw and widely spaced teeth. It was during this period that the species likely attained its highest abundance, and had a relatively continuous distribution. This changed approximately 15,000 years ago with the onset of the current interglacial, which caused the species' Afroalpine habitat to fragment, thus isolating Ethiopian wolf populations from each other.
The Ethiopian wolf is one of five Canis species present in Africa, and is readily distinguishable from jackals by its larger size, relatively longer legs, distinct reddish coat and white markings. John Edward Gray and Glover Morrill Allen originally classified the species under a separate genus, Simenia, and Oscar Neumann considered it to be "only an exaggerated fox". Juliet Clutton-Brock refuted the separate genus in favour of placing the species in the genus Canis, upon noting cranial similarities with the side-striped jackal.
Initial molecular evidence suggested that the Ethiopian wolf is a descendant of the grey wolf, though more recent evidence suggests that this is not the case; although the Ethiopian wolf is closely related to grey wolves, coyotes and golden jackals, it probably diverged some 3-4 million years ago.(Fig. 10)
|Northern Ethiopian wolf|
C. s. simensis
|Rüppell, 1840||Northwest Rift Valley: Simien Mountains, Mount Guna, Guassa Menz, north and south Wollo highlands||crinensis (Erlanger & Neumann, 1900)|
semiensis (Heuglin, 1862)
|Southern Ethiopian wolf|
C. s. citernii
|de Beaux, 1922||Initially classed as a distinct subspecies on account of its bright red coat, though this characteristic is unreliable as a taxonomic distinction. However, its nasal bones are consistently longer than those of the nominate subspecies.||Southeast Rift Valley: Arsi and Bale Mountains|
The Ethiopian wolf is similar in size and build to North America's coyote; it is larger than the golden, black-backed and side-striped jackal, and has relatively longer legs. Its skull is very flat, with a long facial region accounting for 58% of the skull's total length. The ears are broad, pointed and directed forward. The teeth, particularly the premolars, are small and widely spaced. The canine teeth measure 14–22 mm in length, while the carnassials are relatively small. The Ethiopian wolf has eight mammae, of which only six are functional. The front paws have five toes, including a dewclaw, while the hind paws have four. As is typical in the genus Canis, males are larger than females, having 20% greater body mass. Adults measure 841–1,012 millimeters (33.1–39.8 in) in body length, and 530–620 millimeters (21–24 in) in height. Adult males weigh 14.2–19.3 kg (31–43 lb), while females weigh 11.2–14.15 kg (24.7–31.2 lb).
The Ethiopian wolf has short guard hairs and thick underfur, which provides protection at temperatures as low as −15°. Its overall colour is ochre to rusty red, with dense whitish to pale ginger underfur. The fur of the throat, chest and underparts is white, with a distinct white band occurring around the sides of the neck. There is a sharp boundary between the red coat and white marks. The ears are thickly furred on the edges, though naked on the inside. The naked borders of the lips, the gums and palate are black. The lips, a small spot on the cheeks and an ascending crescent below the eyes are white. The thickly furred tail is white underneath, and has a black tip, though, unlike most other canids, there is no dark patch marking the supracaudal gland. It moults during the wet season (August–October), and there is no evident seasonal variation in coat colour, though the contrast between the red coat and white markings increases with age and social rank. Females tend to have paler coats than males. During the breeding season, the female's coat turns yellow, becomes woolier, and the tail turns brownish, losing much of its hair.
Animals resulting from Ethiopian wolf-dog hybridisation tend to be more heavily built than pure wolves, have shorter muzzles and different coat patterns.
Social and territorial behaviours
The Ethiopian wolf is a social animal, which lives in family groups containing up to 20 individuals older than one year, though packs of six wolves are more common. Packs are formed by dispersing males and a few females which, with the exception of the breeding female, are reproductively suppressed. Each pack has a well established hierarchy, with dominance and subordination displays being common. Upon dying, a breeding female can be replaced by a resident daughter, though this increases the risk of inbreeding. Such a risk is sometimes circumvented by multiple paternity and extra-pack matings. The dispersal of wolves from their packs is largely restricted by the scarcity of unoccupied habitat.
These packs live in communal territories, which encompass 6 km2 (2.3 sq mi) of land on average. In areas with little food, the species lives in pairs, sometimes accompanied by pups, and defends larger territories averaging 13.4 km2 (5.2 sq mi). In the absence of disease, Ethiopian wolf territories are largely stable, but packs can expand whenever the opportunity arises, such as when another pack disappears. The size of each territory correlates with the abundance of rodents, the number of wolves in a pack, and the survival of pups. Ethiopian wolves rest together in the open at night, and congregate for greetings and border patrols at dawn, noon and evenings. They may shelter from rain under overhanging rocks and behind boulders. The species never sleeps in dens, and only uses them for nursing pups. When patrolling their territories, Ethiopian wolves regularly scent-mark, and interact aggressively and vocally with other packs. Such confrontations typically end with the retreat of the smaller group.
Reproduction and development
The mating season usually takes place in between August–November. Courtship involves the breeding male following the female closely. The breeding female only accepts the advances of the breeding male, or males from other packs. The gestation period lasts 60–62 days, with pups being born between October–December. Pups are born toothless and with their eyes closed, and are covered in a charcoal grey coat with a buff patch on the chest and abdomen. Litters consist of 2-6 pups, which emerge from their den after three weeks, at which time the dark coat is gradually replaced with the adult colouration. By the age of five weeks, the pups feed on a combination of milk and solid food, and become completely weaned off milk at the age of 10 weeks to six months. All members of the pack contribute to protecting and feeding the pups, with subordinate females sometimes assisting the dominant female by suckling them. Full growth and sexual maturity are attained at the age of two years.
Unlike most social carnivores, the Ethiopian wolf tends to forage and feed on small prey alone. It is most active during the day, the time when rodents are themselves most active, though they have been observed to hunt in groups when targeting mountain nyala calves. Major Percy-Cotton described the hunting behaviour of Ethiopian wolves as thus:
... they are most amusing to watch, when hunting. The rats, which are brown, with short tails, live in big colonies and dart from burrow to burrow, while the cuberow stands motionless till one of them shows, when he makes a pounce for it. If he is unsuccessful, he seems to lose his temper, and starts digging violently ; but this is only lost labour, as the ground is honeycombed with holes, and every rat is yards away before he has thrown up a pawful.
The technique described above is commonly used in hunting big-headed mole-rats, with the level of effort varying from scratching lightly at the hole to totally destroying a set of burrows, leaving metre high earth mounds. Wolves in Bale have been observed to forage among cattle herds, a tactic thought to aid in ambushing rodents out of their holes by using the cattle to hide their presence.
The Ethiopian wolf is restricted to isolated pockets of Afroalpine grasslands and heathlands inhabited by Afroalpine rodents. Its ideal habitat extends from above the tree line at about 3,200m to 4,500m, with some wolves inhabiting the Bale Mountains being present in montane grasslands at 3,000m. Although specimens were collected in Gojjam and northwestern Shoa at 2,500m in the early 20th century, there are no recent records of the species occurring below 3,000m. In modern times, subsistence agriculture, which extends up to 3,700m, has largely restricted the species to the highest peaks.
The Ethiopian wolf utilises all Afroalpine habitats, but has a preference for open areas containing short herbaceous and grassland communities inhabited by rodents, which are most abundant along flat or gently sloping areas with poor drainage and deep soils. Prime wolf habitat in the Bale Mountains consists of short Alchemilla herbs and grasses, with low vegetation cover. Other favourable habitats consist of tussock grasslands, high altitude scrubs rich in Helichrysum, and short grasslands growing in shallow soils. In its northern range, the wolf's habitat is composed of plant communities characterised by a matrix of Festuca tussocks, Euryops bushes and giant lobelias, all of which are favoured by the wolf's rodent prey. Although marginal in importance, the ericaceous moorlands at 3,200-3,600m in Simien may provide a refuge for wolves in highly disturbed areas.
In the Bale Mountains, the Ethiopian wolf's primary prey are big-headed mole-rats, though it will also feed on grass rats, black-clawed brush-furred rats and highland hares. Other secondary prey species include vlei rats, yellow-spotted brush-furred rats, and occasionally goslings and eggs. Ethiopian wolves have twice been observed to feed on rock hyraxes and mountain nyala calves. In areas where the big-headed mole-rat is absent, the smaller East African mole-rat is targeted. In the Simien Mountains, the Ethiopian wolf preys on Abyssinian grass rats. Undigested sedge leaves have occasionally been found in Ethiopian wolf stomachs. It is possible that sedge is ingested for roughage or for parasite control. The species may scavenge on carcasses, but is usually displaced by dogs and golden jackals. It typically poses no threat to livestock, with farmers often leaving herds in wolf inhabited areas unattended.
Range and populations
There are currently six Ethiopian wolf populations. North of the Rift Valley, the species occurs in the Simien Mountains in Gondar, in the northern and southern Wollo highlands, and in Guassa Menz in north Shoa. It has recently become extinct in Gosh Meda in north Shoa and Mount Guna, and have not been reported in Mount Choqa for several decades. Southeast of the Rift Valley, it occurs in the Arsi and Bale Mountains.
|Area||Habitat||Population size estimates||Status||Importance||Threats||Conservation|
|Simien Mountains, North Gondar||Patches connected by corridors, totalling 273 km²||102 (as of 2010)||Stable?||The second largest population, the most genetically diverse, and a tourist attraction.||Human disturbance, road traffic, extensive agriculture, habitat degradation, Helichrysum encroachment into rodent habitat, disease, and competition/predation by golden jackals.||The entire range is within the Simien Mountains National Park, and has been monitored regularly since 2003.|
|Mt. Guna, South Gondar||An isolated patch, estimated at less than 20 km² by 2004||As of 2011, there have been no sightings, despite intensive monitoring during two field visits.||Extinct||Small population, habitat loss, isolation and possible competition with abundant golden jackal populations.||An ORDA Biodiversity Conservation Project is active in the area, in conjunction with woreda and kebele governments, and supported by the Guna Highland Water enterprise.|
|North Wollo Highlands||Patchily distributed in an area of 140 km²||19-23 (as of 2000)||Possibly declining, otherwise stable, as of 1998.||Isolation, habitat degradation, human-wildlife conflict and road encroachment.||The FZS and its associates are working to create the Abuna Yoseph Community Conservation Area, which is to encompass about a third of the wolf's range in North Wollo.|
|South Wollo highlands||Patches connected by corridors, totalling 243 km²||16-19 (as of 2000)||Stable?||After Simien, the second largest area north of the Rift Valley.||Overgrazing, ploughing, persecution and local negative attitudes.||The local ericaceous forests and grasslands are under the protection of the Borena Saiynt Regional Park from agriculture as low as 3,200m. The EWCP and the FZS have been involved in educational programs and wolf monitoring in the Denkoro area.|
|Guassa Menz, North Shoa||A single patch of 112 km²||As of 2010, an estimated 40% have been missing since a CDV outbreak was detected in local dogs.||Although diminishing from disease, the population is healthy and stable.||A core population with ideal habitat, and increasingly a tourist attraction.||Human disturbance, rabies, Helichrysum encroachment into rodent habitat and road traffic.||The wolf's range is protected by community resource management, and the Guassa Community Conservation Area. Educational campaigns are undertaken in schools near wolf ranges.|
|Arsi Mountains, Bale||870 km²||54 wolves in nine packs, as of 2007-2010||Probably declining.||The third largest population, in the second largest Afroalpine area in Ethiopia.||Habitat degradation, expanding agriculture and road traffic.||Protected within the Arsi Mountains Regional Park.|
|Bale Mountains, Bale||1,141 km²||~250 adults and subadults||Declining, but stable in long-term.||The largest population, with the highest density of prey.||Disease (rabies and CDV) and agricultural expansion.||Most of the species' habitat occurs within the Bale Mountains National Park.|
The Ethiopian wolf has been considered rare since it was first recorded scientifically. It is likely that the species has always been confined to Afroalpine habitats, and was thus never widespread. In historical times, all of the Ethiopian wolf's threats are both directly and indirectly human-induced, as the wolf's highland habitat, with its high annual rainfall and rich fertile soils, is ideal for agricultural activities. Its proximate threats include habitat loss and fragmentation (subsistence agriculture, overgrazing, road construction and livestock farming), diseases (primarily rabies and CDV), conflict with humans (poisoning, persecution and road kills), and hybridisation with dogs.
Rabies outbreaks, stemming from infected dogs, have killed many Ethiopian wolves over the 1990s and 2000s. Two well documented outbreaks in Bale, one in 1991 and another in 2008-2009, resulted in the die-off or disappearance of 75% of known animals. Both incidents prompted reactive vaccinations in 2003 and 2008-2009 respectively. CDV is not necessarily fatal to wolves, though there has been a recent increase in infection, with outbreaks of canine distemper having been detected in 2005-2006 in Bale and in 2010 across subpopulations.
During the 1990s, wolf populations in Gosh Meda and Guguftu went extinct. In both cases, the extent of Afroalpine habitat above the limit of agriculture had been reduced to less than 20 km². The EWCP team confirmed the extinction of a wolf population in Mt. Guna in 2011, whose numbers had been in single figures for several years. Habitat loss in the Ethiopian highlands is directly linked to agricultural expansion into Afroalpine areas. In the northern highlands, human density is the among the highest in Africa, with 300 people per km² in some localities, with almost all areas below 3,700m having been converted into barley fields. Suitable areas of land below this limit are under some level of protection, such as Guassa-Menz and the Denkoro Reserve, or within the southern highlands, such as the Arsi and Bale Mountains. The most vulnerable wolf populations to habitat loss are those within relatively low-lying Afroalpine ranges, such as those in Aboi Gara and Delanta in North Wollo.
Some Ethiopian wolf populations, particularly those in North Wollo, show signs of high fragmentation, which is likely to increase with current rates of human expansion. The dangers posed by fragmentation include increased contact with humans, dogs and livestock, and further risk of isolation and inbreeding in wolf populations. Although there is no evidence of inbreeding depression or reduced fitness, the extremely small wolf population sizes, particularly those north of the Rift Valley, raises concerns among conservationists. Elsewhere, the Bale populations are fairly continuous, while those in Simien can still interbreed through habitat corridors.
Encroachment within protected areas
In the Simien Mountains National Park, human and livestock populations are increasing by 2% annually, with further road construction allowing easy access to peasants into wolf home ranges. 3,171 people in 582 households were found to be living in the park and 1,477 outside the park in October 2005. Although the area of the park has since been expanded, further settlement stopped and grazing restricted, effective enforcement may take years. As of 2011, there are about 30,000 people in 30 villages around the park and two within it, including 4,650 cereal farmers, herders, woodcutters and many others. In Bale there are numerous villages in and around the area, comprising over 8,500 households with more than 12,500 dogs. It was estimated in 2007 that the number of households within wolf habitat numbered 1,756. Because of the high number of dogs, the risk of infection in local wolf populations is high. Furthermore, intentional and unintentional brush fires are frequent in the ericaceous moorlands wolves inhabit.
Although wolves in Bale have learned to use cattle to conceal their presence when hunting for rodents, the level of grazing in the area can adversely affect the vegetation available for the wolves' prey. Although no declines in wolf populations related to overgrazing have occurred, it is known that high grazing intensities can lead to soil erosion and vegetation deterioration in Afroalpine areas such as Delanta and Simien.
Human persecution and disturbance
Direct killings of wolves were more frequent during the Ethiopian Civil War, when firearms were more available. The extinction of wolves in Mt. Choqa was likely due to persecution. Although people living close to wolves in modern times believe that wolf populations are recovering, negative attitudes towards the species persist due to livestock predation. Wolves were largely unmolested by humans in Bale, as they were not considered threats to sheep and goats. However, they are perceived as threats to livestock elsewhere, with cases of retaliatory killings occurring in the Arsi Mountains. The Ethiopian wolf has not been recorded to be exploited for its fur, though there was one case where wolf hides were used as saddle pads. It was once hunted by sportsmen, though this is now illegal. Vehicle collisions killed at least four wolves in the Sanetti Plateau since 1988, while two others were left with permanent limps. Similar accidents are a risk in areas where roads cut across wolf habitats, such as in Menz and Arsi.
Hybridisation with dogs
Incidences of Ethiopian wolf-dog hybridisation have been recorded in Bale's Web Valley. At least four hybrids were identified and sterilised in the area. Although hybridisation has not been detected elsewhere, it could pose a threat to the wolf population's genetic integrity, resulting in outbreeding depression or a reduction in fitness, though this does not appear to have taken place.
The Ethiopian wolf is not listed on the CITES Appendices, though it is afforded full official protection under Ethiopia's Wildlife Conservation Regulations of 1974, Schedule VI, with the killing of a wolf carrying a two year jail sentence.
The species is present in several protected areas, including three areas in South Wollo (Bale Mountains National Park, Simien Mountains National Park and Borena Saiynt Regional Park), one in north Shoa (Guassa Community Conservation Area) and one in the Arsi Mountains Regional Park. Areas of suitable wolf habitat have recently increased to 87%, as a result of boundary extensions in Simien and the creation of the Arsi Mountains Regional Park.
Steps taken to insure the survival of the Ethiopian wolf include dog vaccination campaigns in Bale, Menz and Simien, sterilization programs for wolf-dog hybrids in Bale, rabies vaccination of wolves in parts of Bale, community and school education programs in Bale and Wollo, contributing to the running of national parks, and population monitoring and surveying. A 10-year National Action Plan was formed in February 2011.
The species' critical situation was first publicised by the Wildlife Conservation Society in 1983, with the Bale Mountains Research Project being established shortly after. This was followed by a detailed four-year field study, which prompted the IUCN/SSC Canid Specialist Group to produce an action plan in 1997. The plan called for the education of people in wolf-inhabited areas, wolf population monitoring and the stemming of rabies in dog populations. The Ethiopian Wolf Conservation Programme was formed in 1995 by Oxford University, with donours including the Born Free Foundation, Frankfurt Zoological Society (FZS) and the Wildlife Conservation Network.
The overall aim of the EWCP is to protect the wolf's Afroalpine habitat in Bale, and establish additional conservation areas in Menz and Wollo. As well as monitoring wolves in Bale, south and north Wollo, the EWCP carries out education campaigns for people outside the wolf's range in order to improve dog husbandry and manage diseases within and around the park. The program seeks to vaccinate up to 5,000 dogs a year in order to reduce rabies and CDV in wolf inhabited areas.
- This is in reference to the Ethiopian wolf's reported habit of following mares and cows about to give birth in order to feed on the afterbirth.
- Some naturalists ascribe this description to the African wild dog.
- This was later proven incorrect in 2010, when SNP studies showed that the dog's sole ancestor is the grey wolf.
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Marino, J. & Sillero-Zubiri, C. (2011). Canis simensis. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 18 January 2012. Database entry includes a lengthy justification of why this species is endangered
- Sillero-Zubiri, C., and D. Gottelli (1994). Canis simensis. Mammalian Species 385: 1-6.
- Gottelli D, Marino J, Sillero-Zubiri C, Funk S (2004). The effect of the last glacial age on speciation and population genetic structure of the endangered Ethiopian wolf (Canis simensis). Molecular Ecology 13:2275-2286.
- IUCN/SSC Canid Specialist Group 2011, pp. 7–8
- Sillero-Zubiri & MacDonald 1997, p. 5 & 8
- Johnston, Harry Hamilton (1902). The Uganda protectorate; an attempt to give some description of the physical geography, botany, zoology, anthropology, languages and history of the territories under British protection in East Central Africa, between the Congo Free State and the Rift Valley and between the first degree of south latitude and the fifth degree of north latitude. London, Hutchinson & Co. p. 368.
- Lydekker 1908, p. 462
- IUCN/SSC Canid Specialist Group 2011, p. 2
- "The Aberdeen Bestiary". University of Aberdeen. 1995. Retrieved 2012-12-05.
- Smith, C. H. (1839), Dogs, W.H. Lizars, Edinburgh, p. 261
- Rüppell 1835, p. 39
- Powell-Cotton 1902, pp. 206–207
- Parkyns, Mansfield (1853). Life in Abyssinia: Being Notes Collected During Three Years' Residence and Travels in that Country. Vol. II. John Murray. pp. 12-13.
- Darwin, Charles (1868). The Variation of Animals and Plants Under Domestication. Vol. I. Orange Judd. p. 48.
- Vonholdt, B. M.; Pollinger, J. P.; Lohmueller, K. E.; Han, E.; Parker, H. G.; Quignon, P.; Degenhardt, J. D.; Boyko, A. R.; Earl, D. A.; Auton, A.; Reynolds, A.; Bryc, K.; Brisbin, A.; Knowles, J. C.; Mosher, D. S.; Spady, T. C.; Elkahloun, A.; Geffen, E.; Pilot, M.; Jedrzejewski, W.; Greco, C.; Randi, E.; Bannasch, D.; Wilton, A.; Shearman, J.; Musiani, M.; Cargill, M.; Jones, P. G.; Qian, Z.; Huang, W. (2010). "Genome-wide SNP and haplotype analyses reveal a rich history underlying dog domestication". Nature 464 (7290): 898–902. doi:10.1038/nature08837. PMC 3494089. PMID 20237475.
- Dalton, R. 2001. The skull morphology of the Ethiopian wolf (Canis simensis). B.Sc.thesis. University of Edinburgh, Edinburgh, UK.
- Brewer, Douglas J.; Terence, Sir Clark; Phillips, Adrian (March 2002). DOGS IN ANTIQUITY Anubis to Cerberus: The Origins of the Domestic Dog. Aris & Phillips. ISBN 978-0-85668-704-4.
- IUCN/SSC Canid Specialist Group 2011, pp. 2–3
- Powell-Cotton 1902, p. 459
- Clutton-Brock, J., Corbet, G.G., and Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods." Bull. Brit. Mus. Nat. Hist. 29, 119–199.
- Gotelli, D.; C. Sillero-Zubiri, G.D. Applebaum, M.S. Roy, D.J. Girman, J. Garcia-Moreno, E.A. Ostrander, R.K. Wayne (1994). "Molecular genetics of the most endangered canid: the Ethiopian wolf Canis simensis". Molecular Ecology (Institute of Zoology, Zoological Society of London, UK) 3 (4): 301–312. doi:10.1111/j.1365-294X.1994.tb00070.x. PMID 7921357.
- Lindblad-Toh, K.; Wade, CM; Mikkelsen, TS; Karlsson, EK; Jaffe, DB; Kamal, M; Clamp, M; Chang, JL et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature 438 (7069): 803–819. Bibcode:2005Natur.438..803L. doi:10.1038/nature04338. PMID 16341006.
- IUCN/SSC Canid Specialist Group 2011, p. 32
- IUCN/SSC Canid Specialist Group 2011, p. 4
- IUCN/SSC Canid Specialist Group 2011, pp. 4–5
- IUCN/SSC Canid Specialist Group 2011, pp. 3–4
- Powell-Cotton 1902, p. 207
- IUCN/SSC Canid Specialist Group 2011, pp. 19–20
- IUCN/SSC Canid Specialist Group 2011, p. 10
- IUCN/SSC Canid Specialist Group 2011, pp. 40–46
- IUCN/SSC Canid Specialist Group 2011, p. 22
- IUCN/SSC Canid Specialist Group 2011, pp. 22–26
- IUCN/SSC Canid Specialist Group 2011, pp. 26–27
- IUCN/SSC Canid Specialist Group 2011, pp. 27–28
- IUCN/SSC Canid Specialist Group 2011, p. 29
- IUCN/SSC Canid Specialist Group 2011, p. 30
- IUCN/SSC Canid Specialist Group (2011). Strategic Planning for Ethiopian Wolf Conservation. IUCN/SSC Canid Specialist Group. Oxford, United Kingdom.
- Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited.
- Powell-Cotton, P. H. G. (1902). "A sporting trip through Abyssinia : a narrative of a nine months' journey from the plains of the Hawash to the snows of Simien, with a description of the game, from elephant to ibex, and notes on the manners and customs of the natives". London : Rowland Ward.
- Rüppell, Eduard (1835). "Neue Wirbelthiere zu der Fauna von Abyssinien gehörig" (in German). Frankfurt am Main : S. Schmerber.
- Sillero-Zubiri, Claudio; MacDonald, David W. (1997). The Ethiopian Wolf: Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group. ISBN 2-8317-0407-3.
EOL content is automatically assembled from many different content providers. As a result, from time to time you may find pages on EOL that are confusing.
To request an improvement, please leave a comment on the page. Thank you!